Classification of the Cactaceae

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File:Cactaceae serre massey.jpg
A collection of cultivated cacti

In 1984, the International Organization for Succulent Plant Study set up a working party, now called the International Cactaceae Systematics Group, to produce a consensus classification of the cactus family, down to the level of genus. Their classification has been used as the basis for systems published since the mid-1990s. Treatments in the 21st century have generally divided the family into around 125–130 genera and 1,400–1,500 species, which are then arranged in a number of tribes and subfamilies. However, subsequent molecular phylogenetic studies have shown that a very high proportion of the higher taxa (genera, tribes and subfamilies) are not monophyletic, i.e. they do not contain all of the descendants of a common ancestor. As of March 2012, the internal classification of the family Cactaceae is uncertain and likely to change. A possible classification incorporating many of the insights from the molecular studies was produced by Nyffeler and Eggli in 2010.[1]

Introduction

The classification of the family Cactaceae remains uncertain as of March 2012. Since the mid-1990s, the system produced by the International Cactaceae Systematics Group (ICSG) of the International Organization for Succulent Plant Study has been used as the basis of many published classifications. Detailed treatments produced in the 21st century have divided the family into around 125–130 genera and 1,400–1,500 species, which are then arranged into a number of tribes and subfamilies.[2][3][4]

The ICSG classification of the family recognizes four subfamilies: Pereskoideae (consisting only of the genus Pereskia), Opuntioideae, Maihuenioideae (consisting only of the genus Maihuenia) and Cactoideae.[2] Molecular phylogenetic studies suggest that Pereskia is not monophyletic (i.e. its species are not the complete set of descendants of a common ancestor), although the three other subfamilies are.[5][4] The Bayesian consensus cladogram from a 2005 study is shown below:[5]

Cactaceae

 Pereskia Clade A 



 Pereskia Clade B 



Opuntioideae



Maihuenioideae


Cactoideae





Pereskioideae

Five tribes have been recognized within the subfamily Opuntioideae: Tephrocacteae, Pterocacteae, Austrocylindropuntieae, Cylindropuntieae and Opuntieae.[6] All but the first, Tephrocacteae, were shown to be "essentially monophyletic" in molecular phylogenetic study in 2009. A summary of the consensus Bayesian tree is shown below (tribes are bold; the number of species in the study is shown in parentheses).[7]

 Puna (1), Maihueniopsis (3), Cumulopuntia (1) = Tephrocacteae (part)



 Pterocactus (6) = Pterocacteae


terete-stemmed opuntioids

 Maihueniopsis (1), Puna (1), Tephrocactus (7) = Tephrocacteae (part)


 Maihueniopsis (1), Cumulopuntia (5), Puna (1), Austrocylindropuntia (4) ≈ Austrocylindropuntieae


 Quiabentia (1), Pereskiopsis (2), Micropuntia (2), Cylindropuntia (12), Grusonia s.s. (1), Corynopuntia (6) ≈ Cylindropuntieae


flat-stemmed opuntioids

 Miqueliopuntia (1), Tunilla (2), Maihueniopsis (1) 



 Opuntia (2) 


 Brasilopuntia (1), Opuntia (1), Tacinga (2) 


 Opuntia (28), Consolea (6), Nopalea (7) 






≈ Opuntieae

Many of the genera within the Opuntioideae are not monophyletic. Maihueniopsis is highly polyphyletic, appearing in four separate lineages. The two largest genera within the subfamily, Opuntia and Cylindropuntia are also not monophyletic. The classification of the Opuntioideae is thus uncertain as of March 2012; Griffith and Porter say that changes in classification will require "broad information (of multiple data types) regarding all species of opuntioid cacti".[8]

The ICSG classification divides the subfamily Cactoideae into nine tribes. However, phylogenetic research has not supported most of these tribes, nor even the genera of which they are composed. A 2011 study found that "an extraordinarily high proportion of genera" were not monophyletic, including 22 (61%) of the 36 genera in the subfamily Cactoideae sampled in the research.[4] Of the nine tribes recognized within Cactoideae, one, Calymmantheae, comprises a single genus, Calymmanthium.[2] Of the remaining eight, only two (Cacteae and Rhipsalideae) have been shown to be monophyletic. A summary of the cladograms for the Cactoideae presented in a 2011 paper is shown below (ICSG tribes in bold).[9]

Cactoideae

Blossfeldia



Cacteae

core Cactoideae

 Calymmantheae (i.e. Calymmanthium), species of Copiapoa and Frailea


core Cactoideae I

including members of Pachycereeae, Hylocereeae and Browningieae

core Cactoideae II

Rhipsalideae


other core Cactoideae II, including members of Notocacteae, Trichocereeae, Cereeae and Browningieae






The classification of genera and tribes in the Cactaceae is thus likely to change in future.

Classification

Unless otherwise indicated, the subfamily placement of the genera listed here is based on Anderson's 2001 presentation of the ICSG classification, as is the tribal placement of the genera of the subfamily Cactoideae.[10] The division of the subfamily Opuntioideae into tribes is additional to the ICSG system.[11][6]

Subfamily Pereskioideae

Pereskia aculeata
synonyms: Peirescia Zucc. (orth. var.), Peireskia Steud. (orth. var.), Perescia Lem. (orth. var.), Rhodocactus (A.Berger) F.M.Knuth

Subfamily Opuntioideae

Tribe Austrocylindropuntieae

Tribe Cylindropuntieae

including Corynopuntia F.M.Knuth, Micropuntia Daston
synonyms: Peireskiopsis Vaupel (orth. var.)

Tribe Opuntieae

File:Opuntia ficus-indica 4.jpg
Opuntia ficus-indica
synonyms:Airampoa Fric, Cactodendron Bigelow, nom. inval., Cactus Lem., Chaffeyopuntia Fric & Schelle, Clavarioidia Kreuz. (nom. inval.), Ficindica St.-Lag., Nopalea Salm-Dyck, Parviopuntia Soulaire & Marn.-Lap. (nom. inval.), Phyllarthus Neck. ex M.Gómez (nom. inval.), Salmiopuntia Fric (nom. inval.), Subulatopuntia Fric & Schelle, Tunas Lunell, Weberiopuntia Fric
including Nopalea Salm-Dyck

Tribe Pterocacteae

Tribe Tephrocacteae

including Puna R.Kiesling
synonyms: Pseudotephrocactus Fric

Subfamily Maihuenioideae

File:Maihuenia poeppigii 02.jpg
Maihuenia poeppigii

Subfamily Cactoideae

Tribe Browningieae

synonyms: Neocardenasia Backeb.

Tribe Cacteae

synonyms: Anhalonium Lem., Neogomesia Castañeda, Neogomezia Buxb. (orth. var.), Roseocactus A.Berger, Stromatocactus Karw. ex Rümpler (nom. inval.)
File:Astrophytum myriostigma.jpg
Astrophytum myriostigma
synonyms: Astrophyton Lawr. (orth. var.), Maierocactus E.C.Rost
synonyms: Aulacothele Monv. (nom. inval.), Cumarinia Buxb., Glandulifera (Salm-Dyck) Fric, Lepidocoryphantha Backeb., Roseia Fric (nom. inval.)
synonyms: Brittonrosea Speg., Echinofossulocactus Lawr., Homalocephala Britton & Rose
  • Escobaria Britton & Rose – foxtail cactus, pincushion cactus
synonyms: Cochiseia W.H.Earle, Escobesseya Hester, Fobea Fric (nom. inval.), Neobesseya Britton & Rose
synonyms: Bisnaga Orcutt, Brittonia C.A.Armstr. (nom. inval.)
Mammillaria elongata
  • Mammillaria Haw. – fishhook cactus, globe cactus, pincushion cactus, bird's-nest cactus
synonyms: Bartschella Britton & Rose, Cactus L., Chilita Orcutt, Cochemiea (K.Brandegee) Walton, Dolichothele (K.Schum.) Britton & Rose, Ebnerella Buxb., Haagea Fric, Krainzia Backeb., Lactomammillaria Fric (nom. inval.), Leptocladia Buxb., Leptocladodia Buxb., Mamillaria F.Rchb. (orth. var.), Mamillopsis (E.Morren) F.A.C.Weber ex Britton & Rose, Mammariella Shafer (nom. inval.), Mammilaria Torr. & A.Gray (orth. var.), Neomammillaria Britton & Rose, Oehmea Buxb., Phellosperma Britton & Rose, Porfiria Boed., Pseudomammillaria Buxb., Solisia Britton & Rose
synonyms: Napina Fric (nom. inval.), Pseudosolisia Y.Itô (nom. inval.)
synonyms: Navajoa Croizat, Pilocanthus B.W.Benson & Backeb.
synonyms: Encephalocarpus A.Berger
File:Sclerocactus glaucus fh 29 41 COL B.jpg
Sclerocactus glaucus
synonyms: Ancistrocactus Britton & Rose, Coloradoa Boissev. & C.Davidson, Glandulicactus Backeb., Toumeya Britton & Rose
synonyms: Echinofossulocactus Britton & Rose, Efossus Orcutt (orth. var.)
synonyms: Hamatocactus Britton & Rose, Thelomastus Fric (nom. inval.)
synonyms: Gymnocactus Backeb., Normanbokea Kladiwa & Buxb., Rapicactus Buxb. & Oehme

Tribe Calymmantheae

synonyms: Diploperianthium F.Ritter (nom. inval.)

Tribe Cereeae

synonyms: Pierrebraunia Esteves
synonyms: Mirabella F.Ritter, Piptanthocereus (A.Berger) Riccob., Subpilocereus Backeb.
synonyms: Floribunda F.Ritter
synonyms: Buiningia Buxb.
File:Melocactus matanzanus.jpg
Melocactus matanzanus
synonyms: Cactus Britton & Rose
synonyms: Austrocephalocereus Backeb., Siccobaccatus P.J.Braun & Esteves
synonyms: Pilocereus K.Schum., Pseudopilocereus Buxb.

Tribe Hylocereeae

synonyms: Aporocactus Lem., Aporocereus Fric & Kreuz. (orth. var.), Bonifazia Standl. & Steyerm., Chiapasia Britton & Rose, Disisocactus Kunze (orth. var.), Disocereus Fric & Kreuz. (orth. var.), Heliocereus (A.Berger) Britton & Rose, Lobeira Alexander, Mediocereus Fric & Kreuz. (orth. var.), Nopalxochia Britton & Rose, Pseudonopalxochia Backeb., Trochilocactus Linding., Wittia K.Schum., Wittiocactus Rauschert
File:Epiphyllum anguliger (as Phyllocactus darrahii) 91b.jpg
Epiphyllum anguliger
synonyms: Phyllocactus Link, Phyllocereus Miq.
  • Hylocereus (A.Berger) Britton & Rose – nightblooming cactus
synonyms: Wilmattea Britton & Rose
  • Selenicereus (A.Berger) Britton & Rose – moonlight cactus, nightblooming cereus
synonyms: Cryptocereus Alexander, Deamia Britton & Rose, Marniera Backeb., Mediocactus Britton & Rose, Strophocactus Britton & Rose, Strophocereus Fric & Kreuz. (orth. var.)
synonyms: Eccremocactus Britton & Rose, Eccremocereus Fric & Kreuz. (orth. var.), Werckleocereus Britton & Rose

Tribe Notocacteae

synonyms: Pilocopiapoa F.Ritter
File:Eriosyce esmeraldana.jpg
Eriosyce esmeraldana
synonyms: Ceratistes Labour. (nom. inval.), Chileniopsis Backeb., Chileocactus Fric (nom. inval.), Chileorebutia Fric (nom. inval.), Chiliorebutia Fric (orth. var.), Delaetia Backeb., Dracocactus Y.Itô (nom. inval.), Euporteria Kreuz. & Buining, Hildmannia Kreuz. & Buining, Horridocactus Backeb., Islaya Backeb., Neochilenia Backeb. ex Dölz, Neoporteria Britton & Rose, Neoporteria Backeb., Neotanahashia Y.Itô, Nichelia Bullock (nom. inval.), Pyrrhocactus (A.Berger) A.W.Hill, Rodentiophila F.Ritter ex Backeb., Thelocephala Y.Itô
synonyms: Philippicereus Backeb.
synonyms: Acanthocephala Backeb., Brasilicactus Backeb., Brasiliparodia F.Ritter, Brasilocactus Fric (nom. inval.), Chrysocactus Y.Itô (nom. inval.), Dactylanthocactus Y.Itô, Eriocactus Backeb., Eriocephala Backeb., Friesia Fric (nom. inval.), Hickenia Britton & Rose, Malacocarpus Salm-Dyck, Microspermia Fric, Neohickenia Fric, Notocactus (K.Schum.) Fric, Sericocactus Y.Itô, Wigginsia D.M.Porter

Tribe Pachycereeae

synonyms: Dendrocereus Britton & Rose, Monvillea Britton & Rose
synonyms: Bergerocereus Fric & Kreuz. (orth. var.)
File:SaguaroCactusAZ.JPG
Carnegiea gigantea
synonyms: Haseltonia Backeb., Neodawsonia Backeb., Pilocereus Lem.
synonym: Corryocereus Fric & Kreuz. (orth. var.), Erdisia Britton & Rose, Eulychnocactus Backeb. (nom. inval.)
synonyms: Morangaya G.D.Rowley, Wilcoxia Britton & Rose
synonyms: Neoabbottia Britton & Rose
[= Myrtillocactus × Bergerocactus]
synonyms: Myrtillocereus Fric & Kreuz. (orth. var.)
Pachycereus pringlei
synonyms: Pseudomitrocereus Bravo & Buxb., Rooksbya Backeb.
[= Pachycereus × Bergerocactus]
synonyms: Backebergia Bravo, Lemaireocereus Britton & Rose, Lophocereus (A.Berger) Britton & Rose, Marginatocereus (Backeb.) Backeb., Mitrocereus (Backeb.) Backeb., Pterocereus T.MacDoug. & Miranda
synonyms: Cullmannia Distefano, Neoevansia W.T.Marshall, Nyctocereus (A.Berger) Britton & Rose
synonyms: Chichipia Backeb. (nom. inval.), Heliabravoa Backeb.
synonyms: Hertrichocereus Backeb., Isolatocereus Backeb., Isolatocereus (Backeb.) Backeb., Machaerocereus Britton & Rose, Marshallocereus Backeb., Neolemaireocereus Backeb., Rathbunia Britton & Rose, Ritterocereus Backeb.

Tribe Rhipsalideae

File:Rhipsalis salicornioides 1.jpg
Hatiora salicornioides
synonyms: Epiphyllopsis Backeb. & F.M.Knuth, Hariota DC., Pseudozygocactus Backeb., Rhipsalidopsis Britton & Rose
synonyms: Acanthorhipsalis (K.Schum.) Britton & Rose, Acanthorhipsalis Kimnach, Pfeiffera Salm-Dyck
synonyms: Cassytha Mill., Erythrorhipsalis A.Berger, Hariota Adans., Lymanbensonia Kimnach
synonyms: Epiphyllanthus A.Berger, Epiphyllum Pfeiff., Opuntiopsis Knebel (nom. inval.), Zygocactus K.Schum., Zygocereus Fric & Kreuz. (orth. var.)

Tribe Trichocereeae

synonyms: Spinicalycium Fric (nom. inval.)
File:Borzicactus Websteramus, Cleistocactus, Huntington Desert Garden.jpg
Cleistocactus
synonyms: Akersia Buining, Bolivicereus Cárdenas, Borzicactella H.Johnson ex F.Ritter, Borzicactus Riccob, Borzicereus Fric & Kreuz. (orth. var.), Cephalocleistocactus F.Ritter, Cleistocereus Fric & Kreuz. (orth. var.), Clistanthocereus Backeb., Demnosa Fric, Gymnanthocereus Backeb., Hildewintera F.Ritter, Loxanthocereus Backeb., Maritimocereus Akers, Pseudoechinocereus Buining (nom. inval.), Seticereus Backeb., Seticleistocactus Backeb., Winteria F.Ritter, Winterocereus Backeb.
synonyms: Acantholobivia Backeb., Acanthopetalus Y.Itô, Andenea Fric (nom. inval.), Aureilobivia Fric, nom. inval., Chamaecereus Britton & Rose, Chamaelobivia Y. Itô (nom. inval.), Cinnabarinea Fric ex F.Ritter, Echinolobivia Y.Itô (nom. inval.), Echinonyctanthus Lem., Furiolobivia Y.Itô (nom. inval.), Helianthocereus Backeb, Heterolobivia Y.Itô (nom. inval.), Hymenorebulobivia Fric (nom. inval.), Hymenorebutia Fric ex Buining, Leucostele Backeb., Lobirebutia Fric (nom. inval.), Lobivia Britton & Rose, Lobiviopsis Fric (nom. inval.), Megalobivia Y.Itô (nom. inval.), Mesechinopsis Y.Itô, Neolobivia Y.Itô, Pilopsis Y.Itô (nom. inval.), Pseudolobivia (Backeb.) Backeb., Rebulobivia Fric (nom. inval.), Salpingolobivia Y.Itô, Scoparebutia Fric & Kreuz. ex Buining, Setiechinopsis (Backeb.) de Haas, Soehrensia Backeb., Trichocereus (A.Berger) Riccob.
synonyms: Binghamia Britton & Rose, Pseudoespostoa Backeb., Thrixanthocereus Backeb., Vatricania Backeb.
synonyms: Gerocephalus F.Ritter
synonyms: Zehntnerella Britton & Rose
File:Gymnocalycium moserianum.jpg
Gymnocalycium moserianum
synonyms: Brachycalycium Backeb.
synonyms: Floresia Krainz & F.Ritter ex Backeb. (nom. inval.), Haageocactus Backeb. (nom. inval.), Lasiocereus F.Ritter, Neobinghamia Backeb., Peruvocereus Akers
[= Haageocereus × Espostoa]
synonyms: Eriocereus (A.Berger) Riccob., Roseocereus Backeb.
synonyms: Eomatucana F.Ritter
  • Mila Britton & Rose
synonyms: Arequipa Britton & Rose, Arequipiopsis Kreuz. & Buining, Morawetzia Backeb., Submatucana Backeb.
Rebutia flavistyla
synonyms: Aylostera Speg., Bridgesia Backeb., Cylindrorebutia Fric & Kreuz., Digitorebutia Fric & Kreuz., Echinorebutia Fric (nom. inval.), Eurebutia Fric (nom. inval.), Gymnantha Y.Itô, Mediolobivia Backeb., Mediorebutia Fric (nom. inval.), Neogymnantha Y.Itô, Reicheocactus Backeb., Setirebutia Fric & Kreuz. (nom. inval.), Spegazzinia Backeb., Sulcorebutia Backeb., Weingartia Werderm.
synonyms: Meyenia Backeb.

Notes and references

  1. ↑ Lua error in package.lua at line 80: module 'strict' not found.
  2. ↑ 2.0 2.1 2.2 Anderson 2001, pp. 99–103
  3. ↑ Lua error in package.lua at line 80: module 'strict' not found., cited in Bárcenas, Yesson & Hawkins 2011
  4. ↑ 4.0 4.1 4.2 Lua error in package.lua at line 80: module 'strict' not found.
  5. ↑ 5.0 5.1 Lua error in package.lua at line 80: module 'strict' not found.
  6. ↑ 6.0 6.1 Lua error in package.lua at line 80: module 'strict' not found., cited in Griffith & Porter 2009
  7. ↑ Lua error in package.lua at line 80: module 'strict' not found.. Tree based on fig. 1 with tribes from fig. 2.
  8. ↑ Griffith & Porter 2009, p. 114
  9. ↑ Lua error in package.lua at line 80: module 'strict' not found.
  10. ↑ Anderson 2001, pp. 100–103
  11. ↑ Lua error in package.lua at line 80: module 'strict' not found.
  12. ↑ Lua error in package.lua at line 80: module 'strict' not found.

Bibliography

  • Lua error in package.lua at line 80: module 'strict' not found.
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